plants from these hybrids for several generations and eventually obtained

true-breeding, more or less fertile derivatives. In 1930 Meister gave a

botanical description of the new species and named it Triticum seculotricum surutoviense Meister. Lewistsky and Benetzkaja (193 1 ) produced cytological evidence that the new forms produced by Meister from the bread

THE DEVELOPMENT OF TRITICALE

317

wheat X rye crosses amphiploids with 2n = 56 chromosomes. They also

observed univalents and other meiotic irregularities. They assumed that

incompatibility existed among chromosomes between parental genomes.

Since disturbed pairing could not be due to a lack of chromosome homology, they believed that the amphiploids arose as a result of an apogamous

development of F, ovules having a somatic chromosome number and that

this was doubled during the first division of the egg cell.

In his review on early research on triticales, Muntzing (1973a) pointed

out that Lebedeff, working in the Ukraine, produced some rather advanced

cytological work in the early 1930’s. Lebedeff made investigations on

amphiploids that he had created as well as on those of his co-workers.

He suggested that the poor fertility found in the amphiploids was due to

the detrimental effects of inbreeding on the rye genome. This influence

might be avoided by using vigorous self-fertile rye parents. Lebedeff also

demonstrated the formation of unreduced ovules with 28 chromosomes

which produced a 35-chromosome plant when fertilized with rye pollen.

Lebedeffs work on triticale appears to have stopped in the mid 19303,

as did the work at the Saratov Experiment Station under Meister. It was

during this period that Lysenko was gaining a powerful influence in agricultural research in Russia.

Up to this time, triticale was more or less a biological curiosity, as it

appeared to have little or no potential as a commercial crop. Arne Muntzing, at the University of Lund in Sweden, began research on triticale in

1934 and has continued this work to the present time. His contributions

in cytology, genetics, and plant improvement in triticale have been outstanding. His work did much to encourage other scientists to undertake

triticale research. He not only produced new primary octoploid triticales

of his own and secondary forms from crosses between different octoploids,

but also developed a new form in 1933 which was produced by crossing

a tetraploid wheat, Triticum turgidum, with rye and pollinating the F,

hybrid with hexaploid wheat, A triple hybrid with 42 chromosomes resulted, possessing the complete genome of rye and a combination of chromosomes from bread and durum wheat (Muntzing, 1935). Nakajima

(1942) produced triple hybrids in the same way. Shulyndin at Kharkov

produced what he identified as 3-species hybrids, by crossing the F, of

common wheat X rye with durum wheat (Lukyanenko, 1972; Shulyndin,

1972).

Muntzing (1936) observed that occasionally F, plants of wheat-rye hybrids produce anthers having viable pollen. The viable pollen among these

anthers ranged from 20 to 60%. A plant having 56 chromosomes was

produced from controlled self-pollination with this pollen, thus providing

evidence that new amphiploids, which occur occasionally, arise from the

318

F. J. ZILLINSKY

spontaneous formation of small somatic sectors with a doubled chromosome number (Fig. 2 ) . This probably represents the mechanism by which

previous octoploids originated from sterile, haploid hybrids.

Muntzing (1939), reporting on triticale research between 1934 and

1939, observed that crosses between octoploid triticales from different

sources were not as cross-compatible as expected and that fertility of the

hybrids was quite low although the hybrids were vigorous. In later generations more-fertile recombinants could be obtained; these were more productive than the parental strains. He suggested the use of self-fertile ryes

as parents in the production of primary amphiploids to overcome sterility

problems.

B. THE DEVELOPMENT

OF HEXAPLOID

TRITICALE

Two important developments occurred during the late 1930’s that dramatically affected triticale research. First was the discovery that colchicine

could be used to induce chromosome doubling so that new amphiploids

could be produced routinely (Kostoff, 1938). Second, during the same

period improvements in embryo culturing had developed so that hybrids

could be obtained from normally cross-incompatible parental combinations.

These developments paved the way for the production of hexaploid triticales from hybrids between tetraploid wheat and rye (Fig. 3 ) .

The first hexaploid tritical was reported by Derzhavin (1938) from the

cross durum wheat x Secale montanum. A hexaploid triticale from a

durum wheat X cultivated rye, S. cereale, by O’Mara (1948) was to play

an important role in the development of triticale in North America and

Europe. Soon numerous new hexaploid triticales were being produced from

combinations of different tetraploid wheats and diploid ryes (Nakajima,

1952, 1958, 1963; SBnchez-Monge et al., 1956, 1959; Pissarev, 1963;

Kiss, 1966; Larter, 1968; Jenkins, 1969; etc.).

Muntzing (1972) indicated that the first hexaploids to be produced had

such poor seed development that researchers were not encouraged to work

on a form which appeared to have so little economic potential. However,

those produced by O’Mara and Sgnchez-Monge were more promising.

It.

Breeding and Research in Eastern Europe

A.

HUNGARY

Some outstanding work on the development of triticale as a commercial

crop has been done by Arpad Kiss, a Hungarian plant breeder. Kiss started

his investigations on triticale in 1949. Using the species T . turgidum as